Methods in Forest Canopy Research

METHODS IN FOREST CANOPY RESEARCH

METHODS IN FOREST

CANOPY RESEARCH

Margaret (Meg) D. Lowman

Timothy D. Schowalter

Jerry F. Franklin

University of California Press, one of the most distinguished university presses in the United States, enriches lives around the world by advancing scholarship in the humanities, social sciences, and natural sciences. Its activities are supported by the UC Press Foundation and by philanthropic contributions from individuals and institutions. For more information, visit www.ucpress.edu.

University of California Press

Berkeley and Los Angeles, California

University of California Press, Ltd.

London, England

© 2012 by The Regents of the University of California

Library of Congress Cataloging-in-Publication Data

Lowman, Margaret.

Methods in forest canopy research / Margaret D. Lowman, Timothy D.

Schowalter, Jerry F. Franklin.

p.   cm.

Includes bibliographical references and index.

ISBN 978-0-520-27371-9 (cloth : alk. paper)

1. Forest canopies—Research—Methodology. I. Schowalter, Timothy

Duane, 1952– II. Franklin, Jerry F. III. Title.

SD387.F59L69 2012

577.34072—dc23

2012006827

Manufactured in China

19  18  17  16  15  14  13  12

10  9   8   7   6  5   4  3    2   1

The paper used in this publication meets the minimum requirements of

ANSI/NISO Z39.48–1992 (R 2002) (Permanence of Paper).

CONTENTS

Contributors to Text Boxes

Prologue

Preface

Acknowledgments

1. Setting the Stage: Canopy Research Emerges as a Component of Forest Science

2. Forest Types and Site Characteristics

3. Canopy Access Methods: Making It Possible to Study the Upper Reaches of Forests Accurately and Safely

4. Forest Structure and Sampling Units

5. Canopy Conditions, Biota, and Processes

6. Canopy–Atmosphere Interactions

7. Measuring Canopy–Forest Floor Interactions

8. Treetops at Risk? Engaging the Canopy Toolkit in Forest Conservation

9. Conclusions and Recommendations

References

Author Index

Subject Index

CONTRIBUTORS TO TEXT BOXES

YVES BASSET

Smithsonian Tropical Research

Institute, Panama

bassety@si.edu

STEPHANIE A. BOHLMAN

University of Florida

sbohlman@ufl.edu

CATHERINE (CAT) CARDELÚS

Colgate University

ccardelus@colgate.edu

SAMANTHA K. CHAPMAN

Villanova University

samantha.chapman@villanova.edu

BRUNO CORBARA

Global Canopy Programme

b.corbara@globalcanopy.org

SOUBADRA DEVY

Ashoka Trust for Research on Ecology

and Environment (ATREE)

soubadra@atree.org

CHRISTOPHER J. FROST

Warnell School of Forest Resources and

Conservation, University of Georgia

jasmonate@gmail.com (or cfrost@uga.edu)

T. GANESH

Ashoka Trust for Research on Ecology and

Environment (ATREE)

tganesh@atree.org

LEON KAGANOVSKIY

Touro College

leonkag@gmail.com

BEVERLY E. LAW

Department of Forest Ecosystems and

Society, Oregon State University

bev.law@oregonstate.edu

MAURICE LEPONCE

Royal Belgian Institute of Natural Sciences

maurice.leponce@naturalsciences.be

SCOTT E. MILLER

National Museum of Natural History,

Smithsonian Institution

millers@si.edu

WILLIAM R. MILLER

Baker University

william.miller@bakeru.edu

ANDREW MITCHELL

Global Canopy Programme

a.mitchell@globalcanopy.org

VOJTECH NOVOTNY

University of South Bohemia

novotny@entu.cas.cz

GEORGE D. WEIBLEN

University of Minnesota

gweiblen@umn.edu

PHILIP WITTMAN

Tree Foundation

drphil@canopyquest.com

PROLOGUE

The call of timber rang out in the forest as the feller completed his back cut and the immense old-growth Douglas-fir tree began its fall to the ground. Branches, tops, and all matter of organic material and dust filled the air as the tree fell and then burst into hundreds of pieces as it crashed onto the ground. After material had stopped falling, Dr. William Denison, an observer and professor in the Botany and Plant Pathology Department at Oregon State University, began working his way through the debris, shaking his head and despairing of any possibility of reconstructing either the structure or the biological content of the tree’s canopy.

Denison was a key member of the infant research team founded in 1968 to analyze an old-growth Douglas-fir ecosystem at the H. J. Andrews Experimental Forest, deep in the Cascade Range of Oregon. Funding had been provided by the National Science Foundation as a part of the US International Biological Program’s Western Coniferous Forest Biome project. He and Dr. George Carroll, his collaborator at the University of Oregon, had undertaken the task of analyzing branch, foliage, and epiphytic communities of in the canopy of this old-growth forest. Their initial simplistic approach was based on the development of allometric equations, which were needed to estimate such variables, using traditional destructive methods: cut down the trees, sort them into their components, dry and weigh samples, and construct equations. However, this clearly was not going to work with giant trees that ranged up to eighty meters in height and two hundred centimeters in diameter at breast height! Nor did this approach adequately represent the three-dimensional structure of the canopy.

A solution to the dilemma was soon suggested and then proven by two graduate students, Diane Nielsen and Diane Tracy, who were also avid mountain climbers: utilize traditional climbing techniques and equipment to provide repeated nondestructive access to the canopies of these trees. Rigging of many trees followed along with development of booms or spars that provided access to lateral branches and branch-based sampling protocols.

Denison, Carroll, and the rest of their research team went on to develop their pioneering work in temperate canopies of old-growth forests using these techniques and without casualties. Early findings included descriptions of canopy architecture, foliar mass and area, and epiphytic communities in old-growth coniferous forests of the Pacific Northwest. Process-oriented research included the discovery of the important role of nitrogen-fixing canopy lichens, such as Lobaria oregana, in the nitrogen budget of these forests. At about the same time, but independently, similar canopy access methods were being pioneered in tropical forests by Meg Lowman in the Old World tropics and by Don Perry in the New World tropics (see Chapter 1).

This research could be viewed as the beginning of modern canopy studies, crude though it was by current standards. Equipment and techniques for canopy access, instrumentation, and sampling procedures—including the incredible capability existing today for collecting, managing, and analyzing large, spatially explicit data sets—have evolved dramatically in the forty years since that beginning.

Methods for accessing forest canopies have proliferated and continue to evolve. Rope-climbing techniques are highly sophisticated and routine, even in the largest of trees, such as the coast redwood (Sequoia sempervirens) and giant sequoia (Sequoiadendron gigantea). Cranes and walkways provide access to canopies for even the most handicapped of scientists, students, and citizens. Canopy cranes are notable for allowing the use of bulky or heavy instrumentation for in-situ canopy research that would otherwise not be possible. They have also demonstrated their value in allowing access to both the uppermost and outermost extremities of forest canopies and to fragile structures, such as standing dead trees or snags, which are otherwise inaccessible for in-situ research. Many are operational throughout the world, although none currently exist in North America.

Immense progress has occurred in research methodology and instrumentation—and this is what most of this book is about. Perhaps not fully appreciated by current generations of canopy researchers are the incredible advances in hardware and software that have made it possible to organize, store, access, and analyze the huge, spatially explicit data sets that modern instrumentation makes possible—and to do it on personal computers and share it via the web. When Denison and Carroll began their studies, we were still in the era of IBM punch cards and massive centralized computers, where the overnight model run most often yielded a single page reporting formatting error!

Our knowledge of canopy ecosystem structure, function, and composition has expanded exponentially over the last forty years, but we are only at the beginning of our discoveries. Many of those have been unexpected surprises that we sometimes refer to as epiphanies: soils and root systems in tree canopies; old trees that perpetuate crowns and foliage via epicormic branches and shoots; long-lived leaves; the incredible diversity of epiphytes, invertebrates, and even vertebrates rarely or never seen below the canopy; the amazing capacity for water transportation; and many others.

Even so, we are still only at the dawn of our understanding regarding the functioning and biodiversity of the forest canopy universe and their relationships to canopy architecture. An infinitude of discoveries—many unforeseen or even currently unimaginable—remain to be made. Some will involve generalities, so-called principles or theoretical constructs, and others will be specific to the immense diversity of forest types and conditions. All will require the laborious and dedicated efforts of field-oriented scientists, whose efforts this book is intended to serve.

PREFACE

This book is dedicated to the next generation of arbornauts—the technical term for those adventurous scientists who risk gravity, thorns, and stinging ants to explore the higher reaches of forest ecosystems. No work has focused on approaches and techniques for canopy study—although Andrew Mitchell did collate an overview handbook almost ten years ago (Mitchell et al. 2002). Since that time, the field has changed significantly and throughout the world, and forests have become a research priority, especially with the advent of climate change and the understanding of ecosystem services. Issues such as carbon sinks, biodiversity conservation, and canopy–atmosphere interactions have created a plethora of new methods and an urgency to measure the forest canopy accurately. As programs like the National Ecological Observatory Network (NEON; http://www.neoninc.org), the National Science Foundation–funded Long-Term Ecological Research (LTER) sites, and Earthwatch Institute’s multimillion-dollar forest training program for HSBC Bank employees gain momentum and require increasingly labor-intensive methodologies, protocols for sampling become increasingly important to maximize time, funds, and energy.

This book updates the history of approaches to studying forest canopies and surveys the advances in techniques in view of the rapidly changing environmental priorities of the new millennium. Because different questions require a variety of approaches and techniques, this volume is not intended to standardize methods but rather to present advantages and disadvantages of selecting certain approaches and methods for particular objectives. The purpose of writing a book about canopy methods is threefold:

1. Share established methods and recipes for best practices of both observational and experimental techniques for studies of forest canopies.

2. Foster compatibility between different canopy researchers, different forests, and different projects, so that data collected can be compared both within and among research programs to invariably enhance cost-effectiveness, time, and energy.

3. Establish protocols that will lead to more effective forest conservation, using more rigorous data sets that are collected using best practices established by experts in the field.

The authors collectively represent more than one hundred years’ experience in canopy research. Despite this lengthy service to the field, we sought additional experts for all aspects of methods and attempted to tap colleagues who deploy both old as well as new techniques of canopy sampling. The book is intended for a diverse audience, including not only practitioners in the field of forest science but also students of ecology or policy or conservation management, government agencies who wish to understand how field biologists collect and analyze data, citizen scientists, and armchair naturalists and explorers who seek to understand how scientists study and explore forests. Despite our best efforts, this book will not pose solutions to the myriad sampling challenges in forest canopies. For example, the notion of defining vertical transects in forests varies with respect to overall forest structure; in some cases, five meters represents the uppermost canopy, and in other situations, ten meters represents a heavily shaded understory. Similarly, insect sampling may prove exceedingly variable in terms of replication, relative numbers, and both temporal and spatial constraints. The abundance of Diptera may be low in montane forests with relatively cold weather conditions, but they can exceed several thousand per cubic meter in lowland moist tropical forests, making one homogenous sampling methodology difficult to standardize.

In 1982, a slim booklet called Reaching the Rain Forest Roof was published as the outcome of an early canopy symposium at the University of Leeds, England. Just over ten years later, the first canopy textbook, Forest Canopies (Lowman and Nadkarni 1994), was published, with more than 25 chapters, 40 contributing scientists, and 624 pages. During the 1980s, canopy exploration had burgeoned, but as the initial textbook illustrates, it was mainly focused on access and the safety of exploration into what E. O. Wilson affectionately called the eighth continent. Perry (1986) and Lowman (1999) both wrote personal accounts of forest canopy exploration, the latter published in five languages, which opened up canopy research to students worldwide, including K–12 classrooms and citizen scientists, instead of a few privileged researchers in developed countries. Soon after, the second canopy textbook was published, also called Forest Canopies (Lowman and Rinker 2004), totaling 604 pages that focused on hypothesis-driven research. In the last decade, a host of personalized canopy accounts was written (e.g., Lowman et al. 2006; Moffett 1998; Nadkarni 2008; Novotny 2008; Preston 2008, etc.). By providing their stories, these scientists gave citizens, students, and policy makers greater insights into how canopy scientists think and work. By beginning to inform not only scientists but also citizens and decision makers, canopy science has matured into a full-fledged scientific discipline, with its own specialized researchers, techniques, driving questions, and global challenges.

Methods in Forest Canopy Research is organized like a recipe book, where a user can turn to one section without reading the entire book. It does not, however, simply provide specific recipes for exactly how to sample every component of forest canopies: such methodologies are changing constantly and no one method is appropriate for every forest type or for each nuanced hypothesis. Instead, this book offers a basis for how forest canopy methods have developed, why specific methods were designed for different temporal and spatial attributes of forest canopies, and what methods are most appropriate for addressing particular objectives. In sharing the evolution of the last forty years of this arboreal exploration, it exposes the aspects of canopy research that do not yet have adequate methods to address. As soon as this book is printed, new methods will undoubtedly be reported in the literature; this underlies the dynamic and constantly changing elements of current forest canopy research. For this reason alone, we wanted to document the emergence of this new field of forest biology and chart its course for the first forty years of active and dedicated methodology. To this end, we devised a website (http://www.canopymethods.com) to track new techniques and to link to aspects of canopy research, including databases that will ensure improvements in forest data collection. We hope that this virtual element of our canopy methods publication will not only expand the user group of our canopy methods, findings, and hypotheses but also facilitate the necessary updates as the field changes and expands. Over the next forty years, new techniques to document biodiversity; forest processes, including carbon storage; and the growth/mortality/dynamics of the forest, including restoration and impacts of climate change, will take center stage as the next generation of forest biologists tackle the still relatively unknown canopy region.

The first chapter of this book introduces the history of canopy science and why this component of forest research was overlooked until the last few decades. The remaining chapters are structured into sections that reflect different types of canopy research: Chapter 2 measures the canopy structure of different forest types. Chapter 3 describes canopy access methods and how they are executed for specific canopy studies. Chapter 4 surveys biological units within a canopy. Chapter 5 measures the diversity of canopy habitats, biota, and processes. Chapter 6 reviews canopy–atmosphere interactions. Chapter 7 measures canopy–forest floor interactions. Chapter 8 assesses the application of canopy biology to conservation and education. Each chapter has a section of suggested reading and a summary section that highlights the future of canopy research and discusses the ways in which sampling methodologies fall short of adequately addressing important hypotheses. We hope that this book will not only create methods protocols that can be replicated throughout forest canopies around the world but also inspire newer and improved techniques that will ultimately lead to better decision making for forest conservation and management. Throughout the chapters, boxes written by experts highlight canopy methods recipes that are currently considered best practices and/or represent case studies that illustrate creative techniques.

We express enormous gratitude to Dr. Chuck Crumly, editor extraordinaire from University of California Press. Over the years, Chuck has been a major force of inspiration for canopy biology and had an enormous impact on disseminating the field around the world through his professionalism in the publishing world. We also convey our thanks to dozens of colleagues who came out of their treetop eeyries to comment and add new methods to this text. The indomitable spirit and sense of inquiry-based studies by arboreal explorers has given canopy science a unique sense of spirit des corps over the last three critical decades of its development. And last but not least, we thank all the educators and policy makers who have decided that ecosystem services, biodiversity, and forest canopies are worthy of understanding: your work as decision makers to educate the next generation is perhaps more critical than the technical science itself.

ACKNOWLEDGMENTS

The authors extend their arboreal thanks to the international community of canopy scientists who have contributed their sweat, creativity, and inspiration more than four decades of research and exploration. This book would not have seen the light of day without the vision of our editor, Chuck Crumly, who always encourages us to reach for the sky (or at least the treetops). Tim extends his thanks to the Louisiana Agricultural Experiment Station and colleagues associated with the Wind River Canopy Crane Research Facility, Smithsonian Tropical Research Institute, and H. J. Andrews Experimental Forest and Luquillo Experimental Forest Long-Term Ecological Research sites. In addition to those sites listed previously, Meg conveys her appreciation to the North Carolina Museum of Natural Sciences staff and volunteers, in particular mentor and director Betsy Bennett and her amazingly organized assistant Cindy Bogan, without whom this publication would not have ever been completed. She is also grateful to colleagues at North Carolina State University, the Aldo Leopold Leadership Program, ATREE (Ashoka Trust for Research on Ecology and the Environment in Bangalore, India, where much of this was written); Alemayehu Wassie Eshete and the clergy of the Christian Orthodox Church in Ethiopia; and all the other stakeholders in emerging countries who may find this compilation useful to inspire forest conservation from a canopy perspective. Special funds from the TREE (Tree Research, Education, and Exploration) Foundation, Sonia and Larry Ewald, the National Science Foundation, Fulbright, and National Geographic sustained specific components of this book over the years. And finally, Meg thanks her sons Eddie and James for their tree-climbing enthusiasm throughout a childhood of arboreal exploration, whose ancestors would be proud, despite the absence of a tail!

1

SETTING THE STAGE

Canopy Research Emerges as a Component of Forest Science

Indeed, over all the glory there will be a canopy. It will

serve as a pavilion, a shade by day from the heat, and

a refuge and a shelter from the storm and rain.

ISAIAH 4:5–6

Botany needs help from the tropics. Its big

plants will engender big thinking.

E. J. H. CORNER, CAMBRIDGE UNIVERSITY, 1939

THE IMPORTANCE OF FOREST CANOPIES

WHY SUCH A TIME LAG IN STUDYING FOREST CANOPIES?

CANOPY DIVERSITY

FUTURE DIRECTIONS IN CANOPY SCIENCE

THE IMPORTANCE OF FOREST CANOPIES

Our ancestors were tree dwellers. Throughout human history, people have taken to the trees as safe havens, sites of special spiritual connection, and cornucopias for food, medicine, materials, and productivity (reviewed in Lowman 1999; Nadkarni 2008). In many tropical forest regions, indigenous people rely on forests for their livelihoods. Increasingly on a global scale, people and governments are beginning to recognize the importance of ecosystem services provided by forests, which link directly to human health (Perrings et al. 2010). Such benefits include medicine, food, shade, building materials, gas exchange capabilities, energy production, carbon storage, genetic libraries, water cycles, and spiritual/cultural heritage. In many religions, the clergy are stewards of the forests (e.g., Bossart et al. 2006; Cardelús et al. 2012; Lowman 2010; Wassie-Eshete 2007). Children in many cultures climb trees for recreation and build tree houses as an important component of their creative and spiritual links to nature (Louv 2005). With their billions of green leaves that produce sugars from sunlight, the treetops are the engines that support life and the basis of food chains throughout the planet. In an evolutionary sense, humans descended from ancestors in the treetops. Recent findings about ancient hominoids in Ethiopia indicate that human ancestors inhabited forests (not savannahs as was previously thought; White et al. 2009). Anyone who pauses at the zoo to watch a monkey cavorting in tree branches is amused, inspired, and subconsciously reminded of some arboreal sensation that tugs on our evolutionary memory banks.

In Papua New Guinea, a tribe called the Korowai still lives in the treetops, erecting amazing aerial houses accessible by twig ladders. It is speculated that their unusual habit of community tree houses evolved as a mechanism to escape enemies on the forest floor and provide a healthy environment above the dank, dark understory. Tree houses remain a recreational vestige of children and adults alike that inspires links between humans, their ancestry, and the natural world. Many famous people have escaped to childhood tree houses: John Lennon (of the Beatles), Winston Churchill, the Roman Emperor Caligula, and Queen Victoria when she was a young princess (Nelson et al. 2000). Recent medical findings indicate that children who play outdoors and learn about nature have better health and well-being (Louv 2005; Lowman et al. 2009).

Why do the treetops hold such a spiritual and scientific importance for cultures throughout the world? And why have scientists only recently begun to explore these heights for scientific discovery, after decades of studying a mere fraction of the forest understory? Relatively few unknown frontiers of exploration still exist in the twenty-first century, but the treetops are still considered a black box in science (Lowman 1999). The other black boxes for exploration include the ocean floor and soil ecosystems.

Forest canopies reputably house more than 40 percent of the biodiversity of terrestrial ecosystems (reviewed in Lowman and Rinker 2004; Wilson 1992). The combination of sunlight, fruits, flowers, and year-round productivity of the foliage in tropical rain forests provides the ideal conditions for an enormous diversity of inhabitants. Thousands of species of trees and vines produce a veritable salad bar for millions of insects that are in turn eaten by myriad reptiles, amphibians, birds, and mammals, and those primary consumers are eaten by secondary consumers such as harpy eagles, jaguars, and other carnivores. Finally, the cycle of life is completed when soil decomposers break down and recycle all matter for uptake of component nutrients into the canopy (Frost and Hunter 2007).

In addition to classic food chains using energy from sunlight to cycle water and nutrients through leaves to herbivores to carnivores/omnivores to decomposers and back to plants, forests house extra niches for other unique forms of life. Bromeliad tanks, tree crotches, leaf surfaces, and epiphyte communities host extra layers of life in forest canopies. For example, bromeliad tanks house virtual swimming pools in the sky that are home to an entire microcosm of microorganisms. Mosquito larvae, nematodes, tarantulas, katydids, shovel-tailed lizards, and canopy mammals live in and/or drink from these aerial watering holes. Some poison dart frogs trek all the way from the forest floor into emergent trees to deposit their eggs in phytotelms. Other unique canopy niches include the crotches of trees that provide germination sites for strangler figs and soil repositories that house many microarthropods usually associated with the forest floor. Strangler figs are the only trees known to start life at the top and send their aerial roots extending downward, eventually penetrating the soil below to expand and strangle their unwitting host plants. Epiphytes add an extra layer of biodiversity and productivity in the moist, sun-flecked branches. Even more amazing, the surfaces of canopy leaves provide a substrate for epiphylly, another extra layer of plant forms including lichens, mosses, and fungi, many of which grow exclusively on leaf surfaces. Within the canopies of